Car condensatus, from Bouchard (2014).

Belongs within: Cucujiformia.
Contains: Nodostoma, Chrysomelidae, Megalopodidae, Cerambycidae, Belidae, Nemonychidae, Anthribidae, Attelabidae, Brentidae, Brachyceridae, Curculionidae.

The Phytophaga is a major clade of beetles, including the Chrysomelidae (leaf beetles), Cerambycidae (longicorn beetles) and Curculionoidea (weevils). Phytophagans are united by the structure of the pseudotetramerous tarsus, with the fourth tarsal segment minute and concealed between lobes of the third, and of the internal male reproductive organs, with a reduced aedeagus (Grimaldi & Engel 2005). The clade's name reflects the plant-feeding habits of the vast majority of its species; more than 99% of the more than 110,000 described species are plant feeders.

Members of the Curculionoidea have the front of the head extended into an elongate rostrum (secondarily reduced in some subgroups) with a concordant reduction and fusion of many of the mouthparts (such as reduction or loss of the labrum); they also have distinctly clubbed antennae with a long scape lying in a scrobe (groove on the side of the rostrum) (Grimaldi & Engel 2005). In the largest family of weevils, the Curculionidae, the antennae have a distinct elbow, but such an elbow is absent in most members of the other weevil families. Members of the Chrysomeloidea (Chrysomelidae and Cerambycidae) lack a distinct club on the antenna.

The classification of the Curculionoidea has been subject to much debate between authors. For instance, members of the Caridae, a small group of weevils found in Australia and South America, have been variously classified within the Attelabidae, Belidae or Brentidae, and are now generally treated as their own family (Bouchard 2014).

Phytophaga [Bruchelae, Chrysomeliformia, Chrysomelomorpha]
    |--Chrysomeloidea GE05
    |    |  i. s.: Cerambyomima MW15
    |    |--Chrysomelidae GE05
    |    `--+--+--Megalopodidae M05
    |       |  `--Oxypeltidae MW15
    |       |       |--Oxypeltus quadrispinosus B14
    |       |       `--Cheloderus B14
    |       |            |--C. childreni Gray 1832 B14
    |       |            `--C. penai B14
    |       `--+--Cerambycidae MW15
    |          `--+--Anoplodermatinae [Anoplodermatidae] MW15
    |             |    |--Migdolus MW15
    |             |    `--Hypocephalus [Hypocephalini] B14
    |             |         `--H. armatus Desmarest 1832 B14
    |             `--Orsodacnidae MW15
    |                  |  i. s.: Orsodacne C01
    |                  |           |--O. cerasi C01
    |                  |           `--O. ruficollis [=Lema ruficollis] L02
    |                  `--Aulacoscelidinae B14
    |                       |--Janbechynea B14
    |                       `--Aulacoscelis B14
    |                            |--A. appendiculata Cox & Windsor 1999 B14
    |                            `--A. melanocera B14
    `--Curculionoidea [Adelognatha, Cryptoplides, Curculionites, Heteromorphi, Rhynchophora] GE05
         |  i. s.: Episomus pauperatus P66
         |         Etheophanus C88
         |           |--E. striatus C88
         |           `--+--E. optandus C88
         |              `--+--E. nitidellus C88
         |                 `--E. pinguis C88
         |         Dysticheus rotundicollis BM76
         |         Minyomerus languidus BM76
         |         Dyslobus lecontei BM76
         |         Mahakamia kampmeinerti B14
         |         Sciopithes obscurus WZ10
         |         Metamasius quadrilineatus WZ10
         |         Proterhinus O94 [Aglycyderidae W86, Proterhinidae]
         |         Eobelidae W86
         |         Allocorynidae LB91
         |         Cretonanophyes P02 [Eccoptarthridae GE05]
         |           `--C. longirostris P02
         |         Rhina Latreille 1802 L02
         |           |--R. barbicornis [=Curculio barbicornis] L02
         |           `--R. cerasi (Linnaeus 1758) L02, L58 [=Curculio cerasi L02]
         |         Brachyrhinus Latreille 1802 L02
         |           |--B. ligustici (Linnaeus 1758) L02, L58 [=Curculio ligustici L02]
         |           |--B. lineatus (Linnaeus 1758) L02, L58 [=Curculio lineatus L02]
         |           |--B. niger [=Curculio niger] L02
         |           `--B. pyri (Linnaeus 1758) L02, L58 [=Curculio pyri L02]
         |         Campyloscelinae Z93
         |           |--Phaenomerus [Phaenomerini] Z93
         |           `--Tomicoproctus Z93
         |         Aades cultratus (Fabricius 1775) [=Curculio cultratus; incl. C. bicristatus Fabricius 1801] Z93
         |         Alatidotasia exclamationis (Fabricius 1775) [=Curculio exclamationis] Z93
         |         Catoptes Z93
         |           |--C. brevicornis (Broun 1904) (see below for synonymy) E02
         |           `--C. interruptus (Fabricius 1781) [=Curculio interruptus] Z93
         |         Phytoscaphus triangularis [incl. Curculio lanatus Fabricius 1801] Z93
         |         Aoplocnemis clavipes (Fabricius 1775) [=Lixus clavipes] Z93
         |         Cecyropa modesta M94
         |         Mecysmoderes Schoenherr 1837 M94
         |           `--M. stylicornis M94
         |         Peribleptus Schoenherr 1843 M94
         |           |--P. dealbatus (Boisduval 1835) [=Alcides dealbatus, Paepalosomus dealbatus] M94
         |           `--P. scalptus M94
         |--Belidae MW15
         `--+--+--Rhynchitoma MW15
            |  `--+--Nemonychidae MW15
            |     `--Anthribidae MW15
            `--+--Attelabidae MW15
               `--+--+--Brentidae MW15
                  |  `--+--Brachyceridae MW15
                  |     `--Curculionidae MW15
                  `--Caridae [Carinae] MW15
                       |--Cretocar luzzii GE05
                       |--Caenominurus B14
                       `--Car Blackburn 1897 Z94
                            |--*C. condensatus Blackburn 1897 M94
                            |--C. intermedius B14
                            `--C. pini B14

Phytophaga incertae sedis:
  Diapromorpha J89a
    |--D. melanophthalma [=Aspidolopha melanophthalma] J89a
    `--D. melanopus J89a
  Gynandrophthalma J89a
    |--G. divisa Jacoby 1889 J89a
    `--G. sexmaculata Jacoby 1889 J89a
  Melitonoma J89a
    |--M. affine Jacoby 1889 J89a
    `--M. indica Jacoby 1889 J89a
  Chrysolampra J89a
    |--C. curvipes Jacoby 1889 J89a
    `--C. splendens J89a
  Nodostoma J89b
  Aulacolepis decorata J89a
  Demotina J89a
    |--D. balyi Jacoby 1889 J89a
    `--D. serraticollis J89a
  Heteraspis J89b
    |--H. clypeata Jacoby 1889 J89a
    |--H. hebe J89a
    |--H. hirta J89b
    |--H. quadrifasciata Jacoby 1889 J89a
    |--H. variegata Jacoby 1889 J89a
    `--H. vestita J89a
  Acrothinium violaceum Jacoby 1889 J89a
  Corynodes J89a
    |--C. approximans J89a
    |--C. chalybeus J89a
    |--C. fulvicollis Jacoby 1889 J89a
    |--C. gratiosus J89a
    |--C. igneicollis Jacoby 1889 J89a
    |--C. mouhoti J89a
    |--C. peregrinus J89a
    |--C. pretiosus J89a
    `--C. pyrospilotus J89a
  Agasta formosa J89a
  Eumela cyanicollis J89a
  Lygaria westermanni J89a
  Plagiodera marginipennis Jacoby 1889 J89a
  Sphaeroderma J89a
    |--S. biplagiata Jacoby 1889 J89a
    |--S. geminata J89a
    |--S. nigricollis J89a
    |--S. orientalis J89a
    |--S. piceicollis Jacoby 1889 J89a
    `--S. varipes [=Eucycla varipes] J89a
  Argopus J89b
    |--A. angulicollis J89a
    |--A. indicus Jacoby 1889 J89a
    |--A. laevipennis J89b
    `--A. unicolor J89a
  Sphaerophysa piceicollis Jacoby 1889 J89a
  Nisotra bowringi J89a
  Hyphasis J89a
    |--H. feae Jacoby 1889 J89a
    |--H. femoralis Jacoby 1889 J89a
    |--H. indica J89a
    |--H. limbatipennis Jacoby 1889 J89a
    |--H. nigricollis J89a
    |--H. nigricornis J89a
    |--H. parvula J89a
    `--H. unicolor Jacoby 1889 J89a
  Oedionychis birmanica Jacoby 1889 J89a
  Enneamera J89b
    |--E. apicalis Jacoby 1889 J89a
    `--E. sumatrensis J89b
  Pseudodera J89a
    |--P. bifasciata Jacoby 1889 J89a
    `--P. xanthospila J89a
  Amphimela mouhoti J89a
  Clitea picta J89a
  Podontia J89a
    |--P. affinis J89a
    `--P. quattuordecimpunctata J89a
  Ophrida marmorea J89a
  Bonesia J89a
    |--B. balyi Jacoby 1889 J89a
    `--B. clarki J89a
  Morphosphaera sumatrana J89a
  Clitena cyanea J89a
  Hoplasoma J89b
    |--H. ceylonensis J89b
    `--H. ventralis J89a
  Paridea thoracica J89a
  Pseudocophora J89a
    |--P. birmanica Jacoby 1889 J89a
    `--P. buqueti J89a
  Cneorane fulvicornis Jacoby 1889 J89a
  Mimastra J89a
    |--M. chenelli J89a
    |--M. gracilicornis Jacoby 1889 J89a
    `--M. polita Jacoby 1889 J89a
  Luperodes J89b
    |--L. alboplagiatus J89a
    |--L. duodecimmaculata Jacoby 1889 J89a
    |--L. piceicollis Jacoby 1889 J89a
    `--L. scutellatus J89b
  Malaxia viridis [=Glyptolus viridis] J89a
  Sastroides birmanica Jacoby 1889 J89a
  Menippus J89a
    |--M. cervinus J89a
    |--M. cynicus J89a
    `--M. dimidiaticornis Jacoby 1889 J89a
  Haplosonyx J89b
    |--H. nigricollis J89b
    |--H. parvulus J89b
    |--H. scutellata J89a
    `--H. sublaevicollis Jacoby 1889 J89a
  Sphenoraia J89a
    |--S. bicolor J89a
    |--S. duodecimmaculata Jacoby 1889 J89a
    `--S. fulgida J89a
  Leptarthra J89a
    |--L. dohrni J89a
    `--L. fraternalis J89a
  Cynorta melanocephala Jacoby 1889 J89a
  Antipha J89a
    |--A. birmanica Jacoby 1889 J89a
    |--A. bretinghami J89a
    |--A. concolor Jacoby 1889 J89a
    |--A. discoidalis J89a
    |--A. postica J89a
    `--A. puncticollis Jacoby 1889 J89a
  Ochralea J89a
    |--O. divisa Jacoby 1889 J89a
    `--O. nigripes J89a
  Aenidea J89b
    |--A. apicalis Jacoby 1889 J89b
    `--A. sumatrensis J89a
  Syoplia javanensis J89b
  Dorydea J89a
    |--D. basalis Jacoby 1889 J89a
    |--D. indica J89a
    `--D. insignis J89a
  Candezea feae Jacoby 1889 J89a
  Platyxantha gracilis Jacoby 1889 J89a
  Solenia J89a
    |--S. collaris J89a
    `--S. indica Jacoby 1889 [=Euphyma indica] J89a
  Pyropida sumptuosa J89b
  Sphaerometopa acroleuca J89b
  Imolia nigrofasciata J89b
  Lina B66
    |--L. aenea B66
    `--L. formosana Bates 1866 B66
  Adorium chrysomeloides Bates 1866 B66
  Rhinosimus planirostris [=Anthribus planirostris] L02
  Aesernia M86

Catoptes brevicornis (Broun 1904) [=Platyomidia brevicornis non Catoptes brevicornis Sharp 1886 (not preoc. if in dif. gen.)] E02

*Type species of generic name indicated


[B66] Bates, H. W. 1866. On a collection of Coleoptera from Formosa sent home by R. Swinhoe, Esq., H. B. M. Consul, Formosa. Proceedings of the Zoological Society of London 1866: 339–355.

[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[C88] Craw, R. 1988. Continuing the synthesis between panbiogeography, phylogenetic systematics and geology as illustrated by empirical studies on the biogeography of New Zealand and the Chatham Islands. Systematic Zoology 37 (3): 291–310.

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] Band II. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[E02] Emberson, R. M. 2002. The beetle (Coleoptera) fauna of the Chatham Islands: additions and corrections. New Zealand Entomologist 25: 69–77.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[J89a] Jacoby, M. 1889a. Viaggio di Leonardo Fea in Birmania e regioni vicine. XVII.—List of the phytophagous Coleoptera obtained by Signor L. Fea at Burmah and Tenasserim, with descriptions of the new species. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 147–237.

[J89b] Jacoby, M. 1889b. List of the phytophagous Coleoptera collected by Signor Modigliani at Nias and Sumatra, with descriptions of the new species. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 278–287.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[LB91] Lawrence, J. F., & E. B. Britton. 1991. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 543–683. Melbourne University Press: Carlton (Victoria).

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[M86] Macleay, W. 1886. The insects of the Fly River, New Guinea, "Coleoptera" (continued). Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 183–204.

[M94] May, B. M. 1994. An introduction to the immature stages of Australian Curculionoidea. In: Zimmerman, E. C. Australian Weevils (Coleoptera: Curculionoidea) vol. 2. Brentidae, Eurhynchidae, Apionidae and a chapter on immature stages by Brenda May pp. 365–728. CSIRO Australia.

[MW15] McKenna, D. D., A. L. Wild, K. Kanda, C. L. Bellamy, R. G. Beutel, M. S. Caterino, C. W. Farnum, D. C. Hawks, M. A. Ivie, M. L. Jameson, R. A. B. Leschen, A. E. Marvaldi, J. V. McHugh, A. F. Newton, J. A. Robertson, M. K. Thayer, M. F. Whiting, J. F. Lawrence, A. Ślipiński, D. R. Maddison & B. D. Farrell. 2015. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology 40 (4): 835–880.

[O94] Otte, D. 1994. The Crickets of Hawaii: origin, systematics and evolution. The Orthopterists’ Society: The Academy of Natural Sciences of Philadelphia.

[P66] Pascoe, F. P. 1866. Catalogue of longicorn Coleoptera collected in the island of Penang by James Lamb, Esq. Part I. Proceedings of the Zoological Society of London 1866: 222–267.

[P02] Ponomarenko, A. G. 2002. Superorder Scarabaeidea Laicharting, 1781. Order Coleoptera Linné, 1758. The beetles. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 164–176. Kluwer Academic Publishers: Dordrecht.

[WZ10] Wood, D. M., & M. A. Zumbado. 2010. Tachinidae (tachinid flies, parasitic flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 1343–1417. NRC Research Press: Ottawa.

[W86] Wood, S. L. 1986. A reclassification of the genera of Scolytidae (Coleoptera). Great Basin Naturalist Memoirs 10: 1–126.

[Z93] Zimmerman, E. C. 1993. Australian Weevils (Coleoptera: Curculionoidea) vol. 3. Nanophyidae, Rhynchophoridae, Erirhinidae, Curculionidae: Amycterinae, literature consulted. CSIRO Australia.

[Z94] Zimmerman, E. C. 1994. Australian Weevils (Coleoptera: Curculionoidea) vol. 2. Brentidae, Eurhynchidae, Apionidae and a chapter on immature stages by Brenda May. CSIRO Australia.

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